Cannibalistic salamanders, social structures of frogs and pilot whales, and warm-blooded fish were all involved in zoological advances in 1993. In addition, studies of the fossil record challenged traditional theories regarding the origin of avian flight and the ancestry of humans.

Studies on cannibalism in salamanders and on the social structure of whales provided support for theories of kin selection, the tendency to favour genetic relatives over unrelated individuals, by revealing situations in which animals modify their behaviour when they belong to a genetically related family unit. The larval young in some populations of tiger salamanders (Ambystoma tigrinum) are known to become cannibalistic, feeding on other tiger salamanders. The cannibals grow larger than noncannibalistic larvae and develop specialized structures in the mouth that aid in eating other salamanders. Cannibalism occurs most frequently when larvae develop under crowded conditions. David W. Pfennig of Cornell University, Ithaca, N.Y., and James P. Collins of Arizona State University discovered that tiger salamanders reared in genetically unrelated groups are more likely to develop into cannibals than are salamanders raised in groups of siblings. They conducted experiments in which similar-sized larvae were placed in various groups, some being all siblings and some being unrelated. All larvae were of similar size so that variation in body size could not be used by the larvae as a cue to whether individuals were related. The investigators hypothesized that larval salamanders release chemical cues that can be used to distinguish close kin, which have a similar "smell," from unrelated larvae.

Kin-selection theory was also supported by evidence that individual organisms can increase their own genetic success by curtailing breeding and possibly helping their relatives. In a study of the biology of long-finned pilot whales in the Faeroe Islands, southeast of Iceland, Bill Amos of the University of Cambridge and colleagues Christian Schlötterer and Diethard Tautz of the University of Munich, Germany, found that males exhibit atypical behaviour. Pilot whales form large social groups called pods. The investigators used molecular techniques to establish that pod members were closely related, forming an extended family. A pod normally has more adult females than males and may number more than 100 individuals. In most mammalian species in which females live in groups, genetic inbreeding is avoided when male offspring disperse from their homesite before they become breeding adults. Pilot whale males remain with their family pod, yet genetic studies revealed that males in a pod rarely or never breed with the females, who might be their mothers or sisters. Mating is presumably carried out when different pods encounter each other in the ocean. Whether or how the nonbreeding males contribute to the welfare of their relatives in a pod remained to be learned, but defense from marine predators or assistance in a communal feeding effort was suggested. Ironically, the cohesive family structure of long-finned pilot whales makes them prey to human whale hunters. In the early 1990s about 1,700 of the whales were killed each year because pods could easily be herded into coastal areas.

Evidence of the way in which mating systems can develop in the best interest of an individual but not necessarily of the species was presented by Godfrey R. Bourne of Florida Atlantic University in studies of the mating system of a tropical frog, Sinax rubra, in Guyana. If given a choice, females, regardless of body length, select smaller male mating partners, usually about 80% of their own size. Experiments revealed that this size ratio of female to male produced the highest rate of fertilization of a female’s eggs. During mating, a male frog clasps a female and releases sperm while she deposits eggs in the water. A male frog that is larger than the female is not in the proper position for the sperm to reach all of the eggs; thus, he fertilizes significantly fewer. Males smaller than the optimal size do not have enough sperm to fertilize all the eggs. Therefore, to maximize egg fertilization and have the highest reproductive success, a female needs a mate of the proper size and so chooses one accordingly.

A larger male frog, however, often displaces the smaller one that is chosen by the female and ends up mating with her instead. Competition between a small male that is preferred by the female and a large male intruder can reduce female reproductive success because fewer eggs are fertilized. On the other hand, the breeding success of the larger male is enhanced. Smaller males sometimes successfully mate by remaining quiet but alert for approaching females. When a female passes by on her way to check out a calling male, a small, silent "satellite" male may intercept her and mate. This competition among males can reduce the reproductive success of a particular female but ensure the propagation of a particular male’s genes.

Most of the world’s fish species, along with reptiles and amphibians, are ectotherms, or cold-blooded animals, having body temperatures corresponding to that of their surroundings. Endothermic, or warm-blooded, animals have the ability to elevate body temperature internally. The trait is characteristic of mammals and birds as well as some sharks and certain marine fish, including mackerels, tunas, and billfishes (e.g., marlins and swordfish). Two contrasting theories exist to explain what selection pressures were influential in the evolution from ectothermy to endothermy. One theory proposes that endothermy arose following selection for a capability to maintain stable body temperatures across a broad range of environmental temperatures, permitting exploitation of varying thermal conditions. The other proposes that endothermy evolved in response to selection for an increase in aerobic capacity (ability to use oxygen) associated with higher metabolism and a more active lifestyle.

Barbara A. Block and colleagues of the University of Chicago used techniques of molecular genetics to establish the phylogenetic relationships among ectothermic fish species and the three groups of endothermic fishes. They found mackerels, tunas, and billfishes each to be more closely related to ectothermic species than to each other, documenting that endothermy evolved independently in the three different groups. In some species, such as the butterfly mackerel and swordfish, warming is restricted to the central nervous system and retina. The phylogenetic distribution and variable expression of endothermy among the fish groups led the researchers to conclude that endothermy in fishes evolved in response to the advantages of expanding into habitats of varied temperatures, not to a requirement of increased aerobic capacity.

A new living species of large mammal, the first such in more then 50 years, was identified from a physical examination and molecular analysis of skulls, teeth, and skins collected from a largely unexplored rain forest in Vietnam’s mountainous central neck. The animal itself, however, had yet to be seen alive by the scientists involved at the time their findings were published. John MacKinnon of the Asian Bureau for Conservation, Hong Kong, and colleagues of the Vietnamese Ministry of Forestry placed the animal in the Bovidae family, which includes cattle, goats, sheep, and antelopes, and described it as weighing about 100 kg (220 lb) as an adult and having a rich brown coat with white and black markings and sharp, straight horns up to 52 cm (20 in) in length. The new bovid, called the "forest goat" or "spindle horn" by local Vietnamese hunters, was given the name Pseudoryx nghetinhensis.

Two major theories have been proposed for the evolution of flight in birds. One is that flight evolved in ground-dwelling animals that were preadapted for flight. The other is that flight originated in tree-dwelling species. On the basis of fossils from the Late Jurassic (150 million years ago), Archaeopteryx is the generally accepted predecessor of flying birds and the focus of most theoretical discussions on the origin of avian flight. During the year Alan Feduccia of the University of North Carolina lent credence to the origin of flight from tree-dwelling forms by means of a study that compared the claw geometry of Archaeopteryx with that of modern birds. The claw curvature of the ancient bird was shown to be similar to that of modern birds that perch in trees or climb tree trunks rather than to that of ground-dwelling birds, suggesting that this earliest known feathered ancestor of birds was arboreal. (See Ornithology, below.)

Among the most controversial evolutionary interpretations from the fossil record are those surrounding the relationships between humans and other primates. David R. Begun of the University of Toronto examined fossil hominids from Hungary estimated to be 10 million years old. He concluded that they may be the closest known relatives of chimpanzees, gorillas, and humans. In addition, his findings supported the view that humans are more closely related evolutionarily to chimpanzees than either are to gorillas, a position held by many molecular biologists.

Entomology

The fossil record of Insecta, the most diverse class of living animals, has received less attention in the English-language scientific literature than many other major animal groups. Part of the reason lies with the perception that insect fossils are rare. Refuting that notion, a study by Conrad C. Labandeira of the Smithsonian Institution, Washington, D.C., and J. John Sepkoski, Jr., of the University of Chicago revealed that insect diversity has exceeded that of four-limbed vertebrates since Carboniferous times (about 325 million years ago). The investigators compiled geochronological records for 1,263 insect families, relying on extensive fossil records reported in German, Russian, and Chinese literature. Objectives of the study were to determine the fossil diversity and rates of evolution of insects and to relate these data to the worldwide development of angiosperms (flowering plants) that originated in the Cretaceous Period (about 125 million years ago). One conclusion was that the high diversity and radiation of modern insect families began nearly 100 million years before flowering plants first appeared, rather than after and in response to their appearance. The researchers also concluded that the increasing diversity of insect families has persisted over geologic time because of low extinction rates rather than because of high rates of evolution during particular periods.

The merit in using fossilized material to interpret evolutionary relationships is often controversial. A study reported during the year, however, demonstrated the utility of using extinct insects to resolve a dilemma regarding the relationships between major groups of organisms. An Australian termite, Mastotermes darwiniensis (order Isoptera), had long been considered the most primitive isopteran and the "missing link" between cockroaches and termites. To establish the relationship between termites and roaches, Rob DeSalle, Ward Wheeler, and David Grimaldi of the American Museum of Natural History, New York City, and John Gatesy of Yale University used molecular techniques to examine and compare DNA sequences from the genes of M. darwiniensis and other living species of insects as well as an extinct termite (M. electrodominicus) from the Dominican Republic. The fossil, preserved in amber 25 million-30 million years old, yielded what was at the time the oldest DNA extracted from a fossil. (In mid-1993 scientists reported recovering DNA from a weevil encased in amber 120 million-135 million years old.) The investigators concluded that termites, including the genus Mastotermes, are a monophyletic group (all derived from the same common ancestor) that evolved independently from the roaches.

Most flies (order Diptera) emit and hear low-frequency (100-500 Hz) sounds that travel short distances, whereas crickets emit high-frequency (usually above three kilohertz) sounds audible at much greater distances. Daniel Robert and Ronald R. Hoy of Cornell University and John Amoroso of the University of Florida reported the discovery of a parasitoid fly (genus Ormia) having an ear capable of detecting high-frequency sounds made by crickets (genus Gryllus). Male field crickets produce far-reaching high-frequency sounds to attract females; however, female parasitoid flies are also attracted to the calling males, on or near which they deposit larvae that burrow into the host cricket. The cricket dies within 10 days, by which time the larvae have developed into pupae that emerge. The newly discovered hearing organ (tympanic ear) in the fly is anatomically and functionally characteristic of a cricket’s and represents an instance of convergent evolution that allows the fly to exploit the mating behaviour of its host.

James T. Cronin and Donald R. Strong of the University of California at Davis conducted experiments to examine egg-laying patterns of a parasitoid, the fairyfly wasp (Anagrus delicatus), in relationship to its plant hopper host, Prokelisia marginata. Plant hoppers, the most abundant herbivorous insects in the Atlantic and Gulf coastal marshes of North America, both feed and lay eggs on salt marsh cord grass (Spartina alterniflora). The female wasp seeks out and lays its eggs only in the eggs of plant hoppers. The investigators measured the time the wasps took to search grass leaves for plant hopper eggs and then to deposit their eggs. They discovered that the wasps spent more than an hour on a plant once plant hopper eggs had been located. Although other plant hopper eggs were available to parasitize, a female wasp laid only a few of her eggs before leaving to search other plants, thus distributing her eggs among different leaves of grass. Such behaviour stood in contrast to the traditional view that parasitoids minimize the time invested in egg-laying activity. The researchers found that 20-30% of the cord grass leaves in the habitat aged and died during the approximately 24 days required for parasitoid larval development, resulting in deaths of the eggs of both species of insects. One conclusion was that, although wasp egg-laying rates are lower than can be achieved, the strategy of spreading eggs among several grass patches increases the probability that at least some offspring survive.

Ornithology

The ability of captive African gray parrots (Psittacus erithacus) to mimic human speech and other sounds is well known, but observations of wild populations in West Africa had not indicated that they practice vocal mimicry naturally; that is, of the kind commonly seen in such birds as mockingbirds and starlings. However, analysis of a sound recording of a gray parrot in Zaire revealed the unmistakable reproduction of sounds from nine bird species and one kind of bat, the first evidence of sound copying by gray parrots in the wild. Furthermore, additional tapes of wild gray parrots in Gabon and Côte d’Ivoire suggested that such impressionism may be widespread.

That American blue jays eat large numbers of acorns in autumn and bury many more for winter consumption has long interested ornithologists, for although these nuts contain unpalatable tannins known to upset the digestive enzymes of other animals that consume them, blue jays appear to suffer no harm. Carter Johnson of South Dakota State University discovered that jays eating acorns that had been invaded by acorn weevil larvae suffered no weight loss provided that each bird consumed with the nuts roughly 100 larvae a day. By comparison, other jays that ate only pristine, uninfested nuts did lose weight.

An individual Clark’s nutcracker, another hoarder, may hide 30,000 conifer seeds in 6,000 separate holes in the forest floor. The birds successfully retrieve many of the seeds, displaying an excellent spatial memory, but Alan Kamil of the University of Massachusetts at Amherst and J.P. Balda of Northern Arizona University found in experimenting with some birds’ orienting ability that the birds relocated seeds whether or not they approached each cache from the same direction as when burying the seeds. Thus, instead of relying on direction as an aid to memory, the nutcrackers may generate a kind of "cognitive map."

In the U.K., where the breeding biology of common bird species probably has been more widely studied than in other countries, egg-laying dates for 33 species of the 82 studied showed a trend, over the 30 years to 1990, toward earlier laying. Among the species the advance varied from one to 22 days, with a mean of 8 days. One contributory cause could be global warming.

The dunnock, or hedge sparrow, a small dun-coloured European perching bird (passerine), was the subject of a 10-year study by N.B. Davies of the University of Cambridge published as Dunnock Behaviour and Social Evolution. Within a population of dunnocks, nearly every conceivable mating system can be found. Some males monopolize the sexual favours of two females, while others have but one mate and still others share either one or two females with another male. Why then, when most birds are monogamous or nearly so, do dunnocks have such a variable mating system?

In broad outline Davies’ finding is that the dunnock’s mating system is a product both of a variable ecology and of conflict between individuals. A female will defend a territory large enough to satisfy her nutritional requirements. Males then defend territories that enclose female territories and, in so doing, control the reproductive opportunities of the females. Some territories are so large, however, as to require two males to defend the one or two occupant females. Thus commences one of the many conflicts. A female prefers both males to mate with her so that both stay and feed the offspring. In contrast, each male prefers to monopolize the female. Thus, the dominant male attempts to guard the female and keep his weaker rival at bay. The female, preferring the attention of both males, attempts to find the weaker male, who will be enticing her at a distance from a bush. It is the female’s task to elude the dominant male. Once she has done so, he will flit about frantically looking for the pair so as to break them up.

Knowledge of Archaeopteryx, the most well-known ancient fossil bird, comes from a half-dozen specimens found in Bavarian rocks about 150 million years old. A somewhat younger fossil bird, Sinornis satensis, which dates from about 135 million years ago, had been known from only two specimens, one from Spain and one from Mongolia, until a third, more complete specimen, found in China, was described in the early 1990s. It was the only one of the three to be found with intact hand bones, which reveal the transition from reptilian forelimb to avian flight wing. Furthermore, the specimen displays a grooved wrist bone, which would have enabled this early bird to fold its wing back as modern birds can. On the other hand, Sinornis also shows a short, toothed reptilian snout and a lizardlike pelvis. In 1993 a still younger fossil bird, 75 million years old, was described from two partial skeletons unearthed in Mongolia. The species was flightless, having had stubby arms ending in a large single claw, and may have evolved from an earlier flying form, as did rheas, emus, and ostriches. Named Mononychus olecranus, meaning "one claw, elbow head," the fossil bird appeared more closely related to modern birds than to Archaeopteryx. (See Zoology, above.)

An average of two to three fully scientifically defensible discoveries of new bird species are made each year, adding to the approximately 9,250 living species known. In the past two years new discoveries included two warblers from China: the Chinese leaf warbler (Phylloscopus sichuanensis), distinguished from its closest relative, P. chloronotus, by its very different song and calls, and the Hainan leaf warbler (P. hainanus), a distinctively deep-yellow species.

A summary of the results of an exceptionally long-term study (more than 40 years) of the fulmar, a seagoing petrel, revealed that males most commonly do not first breed until they are 10 years old and females, 12 years. The fulmar’s mean adult life span appeared to be about 34 years, and the oldest known individual died at about age 46.